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SHORT COMMUNICATION Identifying Copepod Functional Groups From Journal of Plankton Research plankt.oxfordjournals.org J. Plankton Res. (2016) 38(1): 159–166. First published online November 3, 2015 doi:10.1093/plankt/fbv096 SHORT COMMUNICATION Identifying copepod functional groups Downloaded from from species functional traits FABIO BENEDETTI, STE´ PHANE GASPARINI AND SAKINA-DOROTHE´ EAYATA* http://plankt.oxfordjournals.org/ SORBONNE UNIVERSITE´ S, UPMC UNIV PARIS 06, INSU-CNRS, LABORATOIRE D’OCE´ ANOGRAPHIE DE VILLEFRANCHE, 181 CHEMIN DU LAZARET, 06230 VILLEFRANCHE-SUR-MER, FRANCE *CORRESPONDING AUTHOR: [email protected] Received June 15, 2015; accepted October 13, 2015 Corresponding editor: Roger Harris by guest on March 22, 2016 We gathered information on the functional traits of the most representative copepod species in the Mediterranean Sea. Our database includes 191 species described by 7 traits encompassing diverse ecological functions: minimal and maximal body length, trophic group, feeding type, spawning strategy, diel vertical migration and vertical habitat. Cluster analysis in the functional trait space revealed that Mediterranean copepods can be separated into groups with distinct ecological roles. KEYWORDS: copepods; functional trait; zooplankton; Mediterranean Sea Functional traits are phenotypic characteristics of organ- functional traits for zooplankton taxa. However, these isms that impact their fitness and are relevant to ecosys- functional traits have mainly been documented from la- tem function (Violle et al., 2007). For zooplankton, traits boratory experiments, thereby only encompassing a small can be classified according to ecological functions— number of species (Barnett et al., 2007; Saiz and Calbet, feeding, growth/reproduction, survival— and types— 2007; Barton et al., 2013; Kiørboe et al., 2015). For appli- morphological, physiological, behavioural, life history cation in marine ecology, traits are needed at the species (Litchman et al., 2013). As organisms have to allocate level and for the largest possible number of species, energy among ecological functions, traits relate to one which requires the gathering of a huge amount of specific another through trade-offs (Kiørboe et al., 2015). information. Such a challenge should be tackled step by Litchman et al. (Litchman et al., 2013) recently advocated step, focusing on some taxa and/or ocean basins at first. the implementation of a comprehensive matrix of key Functional traits could be used to gather species with available online at www.plankt.oxfordjournals.org # The Author 2015. Published by Oxford University Press. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited. JOURNAL OF PLANKTON RESEARCH j VOLUME 38 j NUMBER 1 j PAGES 159–166 j 2016 similar traits into functional groups (i.e. sets of species All body sizes were obtained from Razouls et al. with similar effects on ecosystem functions and/or (Razouls et al., 2005–2015), whereas the other traits were similar response to environmental conditions; Gitay and obtained from an extensive literature review (see Noble, 1997) or to describe functional diversity of zoo- Supplementary data, Material S4 and Table SII for the plankton communities (Barnett et al., 2007; Vogt et al., full list of references). Additionally, to discuss the potential 2013; Pomerleau et al., 2015). Identifying and describing role of each functional group in the pelagic ecosystem, zooplankton functional groups should then increase our the species’ preferential depth layer was established understanding of zooplankton ecological roles in marine (epi-/meso-/bathypelagic). We were able to determine ecosystems. at least 5 of the 7 functional traits for 171 species. The In this study, we developed a trait database for the most trait database for the 191 copepod species is available as commonly sampled and abundant Mediterranean copepod Supplementary Table SII and can also be downloaded species. Copepods are abundant in marine pelagic ecosys- from PANGAEA (http://doi.pangaea.de/10.1594/ tems (e.g. Siokou-Frangou et al., 2010) where they constitute PANGAEA.854331). the main trophic link between primary producers and In order to identify functional groups, we performed a higher trophic levels, such as small pelagic fishes (Costalago multiple correspondence analysis (MCA) on the trait et al., 2015). Copepods are also relatively well documented matrix. MCA is an ordination method in reduced space Downloaded from in terms of distribution and biology (Razouls et al., 2005– for the multivariate analysis of categorical variables 2015). From this trait database, the aim of this study was to (Husson et al., 2010). The computation of the MCA func- estimate how many functional groups could be identified tional space was performed on four traits: class of among Mediterranean copepods, to characterize them maximum body length (Size_1: 0.501.80 mm, Size_2: and to discuss their ecological significance. 1.89–2.85 mm, Size_3: 3.00–5.70 mm, Size_4: 6.10– http://plankt.oxfordjournals.org/ We considered 191 copepod species (Table I) that are 11.0 mm), binary trophic group (Carnivore, Omnivore, the most representative of the Mediterranean copepod Herbivore, Detritivore), feeding type and spawning strat- communities, in terms of both abundance and presence egy. Indeed, the minimum body length was highly corre- (Siokou-Frangou et al., 2010; Mazzocchi et al., 2014). lated to the maximum body length (R2 ¼ 0.866, n ¼ 191) More details on how this list of species has been gathered and DVM behaviour was not taken into account since it are available in Supplementary data, Material S1. We tends to be very plastic for most species, meaning that it used the following traits covering various types and eco- is known to vary greatly according to the environmental logical functions (Litchman et al., 2013), known to be eco- fluctuations and species’ ontogeny (see Pomerleau et al., logically meaningful (Kiørboe and Sabatini, 1994; 2015). A preliminary MCA incorporating these traits by guest on March 22, 2016 Kiørboe, 2011; Kiørboe et al., 2015) and commonly used showed they had no impact in the definition of functional for zooplankton (Barnett et al., 2007; Barton et al., 2013; groups. Species for which the four traits were not fully Pomerleau et al., 2015): defined were used as supplementary objects, meaning that they are associated with a group a posteriori,from - two morphological traits which relate to many eco- their informed traits (see Supplementary data, Material logical traits: minimum and maximum adult body S2 for more details on the MCA). The Euclidean distance (cephalothorax) length (mm); among the 191 species in the functional space was com- - one physiological trait defining the species’ trophic puted using their coordinates along the four significant group (Carnivore, Omnivore–Carnivore, Omnivore, axes of the MCA (70.77% of the variance). Hierarchical Omnivore–Herbivore, Omnivore–Detritivore); agglomerative clustering was performed on this distance - one behavioural trait depicting feeding strategy,classified matrix using a synoptic aggregation method (Ward’s; into three classes (Kiørboe, 2011): active ambush Husson et al., 2010). Depending on the cutting level, two, feeding, cruise feeding and filter feeding, mixed feeding three or six clusters could be identified (Fig. 1). The first (for species that can switch between the three strategies); level distinguished species according to the trophic - one life history trait related to reproduction, defining the group (carnivore vs. non-carnivore). Among non-carnivore egg-spawning strategy (broadcast-spawner, sac-spawner); species, the second level separated broadcasters from sac- - one behavioural trait related to diel vertical migration spawners. Then, each of these groups was divided into (DVM) behaviour, classified into four classes accord- two subgroups with different size and/or feeding type. ing to the intensity of the observed migration: Since higher cut levels could not be clearly related to func- Non-migrant, Weak migrant (DVM occurs within tional traits, six functional groups were retained (Tables I tens of metres), Strong migrant (over several hundreds and II). We will now detail each functional group and of metres), Reverse migrant (for species that migrate discuss their ecological role in the Mediterranean pelagic deeper at night). food web. 160 Table I: List of the 191 Mediterranean copepod species whose traits have been described Functional group 123 4 56 Number Species Number Species Number Species Number Species Number Species Number Species F.BENEDETTI 5 Aetideopsis armata 66 Aetideus armatus 11 Acartia clausi 8 Anomalocera 69 Diaixis pygmaea 45 45 Clausocalanus patersoni arcuicornis 10 Augaptilus 77 Aetideus 22 Acartia danae 12 12 Calanoides carinatus 70 Disco minutus 46 46 Clausocalanus longicaudatus giesbrechti furcatus 11 Augaptilus spinifrons 29 Candacia bispinosa 33 Acartia discaudata 13 Calanopia elliptica 71 Distioculus minor 47 47 Clausocalanus jobei 27 Candacia armata 31 Candacia 44 Acartia negligens 14 14 Calanus helgolandicus 96 Homeognathia 50 50 Clausocalanus ET AL giesbrechti brevis parapergens 28 Candacia bipinnata 34 Candacia simplex 9 Archescolecithrix 15 Calocalanus adriaticus 113 Monothula subtilis 51 51 Clausocalanus . auropecten paululus j COPEPOD FUNCTIONAL GROUPS FROM SPECIES FUNCTIONALTRAITS 30 Candacia ethiopica
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